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The Origin of Species AP Chapter 24 Microevolution vs macroevolution • Microevolution – changes in gene frequencies within population • Macroevolution – origin of new taxonomic groups, ie – species level and above Speciation is at the boundary between microevolution and macroevolution. What is a species? •Species is a Latin word meaning “kind” or “appearance.” •Traditionally, morphological differences have been used to distinguish species. •Today, differences in body function, biochemistry, behavior, and genetic makeup are also used to differentiate species. 4 approaches to species concept 1. Biological – reproductive isolation 2. Morphological – anatomical differences 3. Ecological – unique roles in environment 4. Phylogenetic – based on evolutionary lineage with distinct morphology and molecular sequences Mayr’s concept of species • 1942 • Potential to interbreed and produce fertile offspring The biological species concept is based on interfertility rather than physical similarity Figure 24.2a Figure 24.2b The biological species concept is based on interfertility rather than physical similarity Interfertility concept does not apply to • Asexually reproducing organisms • Extinct species Barriers to speciation • Prezygotic – prevent mating or successful fertilization • Postzygotic - prevent the hybrid zygote from developing into a viable, fertile adult. Prezygotic barriers • Habitat isolation • Temporal isolation – different breeding times • Behavioral isolation • Mechanical isolation • Gametic isolation Fig. 24-4h (f) Mechanical Isolation Bradybaena with shells spiraling in opposite directions Fig. 24-4e (c) Temporal Isolation Eastern spotted skunk (Spilogale putorius) Western spotted skunk (Spilogale gracilis) These skunks have different breeding times. Fig. 24-4g (e) Behavioral Isolation Courtship ritual of bluefooted boobies Fig. 24-4k (g) Gametic isolation in sea urchins Sea urchins – Their gametes will not fuse with other gametes of other species. Postzygotic Barriers •Reduced hybrid viability - Genetic incompatibility between the two species may abort the development of the hybrid at some embryonic stage or produce frail offspring. •Reduced hybrid fertility - Even if the hybrid offspring are vigorous, the hybrids may be infertile and the hybrid cannot backbreed with either parental species. Reduced hybrid breakdown – In some cases, first generation hybrids are viable and fertile.However, when they mate with either parent species or with each other, the next generation is feeble or sterile. Fig. 24-4m Reduced Hybrid Fertility (i) Mules are infertile. Modes of speciation 1) Allopatric – geographical separation Figure 24.7 Allopatric speciation of squirrels in the Grand Canyon Modes of speciation 2. Sympatric – biological barriers prevent gene flow in overlapping populations as in autopolyploidy, allopolyploidy, mate preference, etc. Figure 24.8 Has speciation occurred during geographic isolation? Examples of Sympatric Speciation involving polyploidy (extra sets of chromosomes) which can lead to new species. • Autopolyploidy – more than two sets of chromosomes; meiotic failure of chromosomes to separate, common in selfpollination in plants • Allopolyploidy * – interspecific hybrid; may become fertile due to nondisjunction in formation of gametes • * more common Sympatric speciation by autopolyploidy in plants Notice that the chrom do not separate! Figure 24.15 One mechanism for allopolyploid speciation in plants “allo” means coming from another place, in this case another species. Fig. 24-UN2 Ancestral species: AA Triticum monococcum (2n = 14) BB Wild Triticum (2n = 14) Product: AA BB DD T. aestivum (bread wheat) (2n = 42) DD Wild T. tauschii (2n = 14) • Around 1870, a new species of grass turned up at the salt marches near the coast of the English Channel: Spartina townsendii . It was taller than the indigenous Spartina alternifolia. • Another relative, Spartina stricta, inhabits the NorthAmerican east coast. It was brought in to Europe and began to occupy the sites of Spartina alternifolia. • It was now suspected that Spartina townsendii was a hybrid of the two original species. The fact that Spartina townsendii has 2n = 126 chromosomes, Spartina alternifolia has 2n = 70 and Spartina stricta has 2n = 56 chromosomes makes this suggestion seem likely. Modes of Speciation 3. Parapatric Speciation • Involves both time and space • Is speciation at the perimeter of the ancestral species range where the environment changes in a qualitative way • Local environment or resources available at the margin of the species range are sufficiently different that natural selection selects for different adaptations • Natural selection would be against the hybrids Hybrid zones • Where divergent allopatric and parapatric populations come back and interbreed Three outcomes… With renewed or continued contact between two populations, there are three possible outcomes: 1. Individuals can hybridize readily. No speciation 2. Individuals do not hybridize at all (reinforcement) Full speciation 3. Individuals hybridize but offspring have reduced fitness. Speciation in progress. Selection for evolution of strong reproductive barriers. 32 Fig. 24-14-4 Strengthening of reproductive barriers Isolated population diverges Possible outcomes: Hybrid zone Reinforcement OR Fusion Gene flow Hybrid OR Barrier to is great If gene flow gene flow Population enough, the parent species Stability (five individuals can fuse into a single are shown) species Hybrids continue to be produced between the two species in the area of their overlap, but the gene pools of both parent species remain distinct. Fig. 24-15 Sympatric male pied flycatcher 28 Allopatric male pied flycatcher Pied flycatchers 24 Collared flycatchers Number of females Reinforcement of barriers to reproduction, has to be strong in sympatric species 20 16 12 8 4 (none) 0 Females mating Own Other with males from: species species Sympatric males Own Other species species Allopatric males Fig. 24-16 Pundamilia nyererei Pundamilia pundamilia Gene pools have fused. Pundamilia “turbid water,” hybrid offspring from a location with turbid water Timing of evolution? • Gradual (Lyell, Darwin) • Punctuated equilibrium (Gould) – evolution occurs in spurts and interspersed within long periods of stasis Species undergo most morphological modifications when they first bud from their parent population. After establishing themselves as separate species, they remain static for the vast majority of their existence. Anagenesis vs cladogenesis Speciation Rates • The punctuated pattern in the fossil record and evidence from lab studies suggests that speciation can be rapid • The interval between speciation events can range from 4,000 years (some cichlids) to 40,000,000 years (some beetles), with an average of 6,500,000 years Case Study As the Worm Turns: Are apple maggot flies and hawthorn maggot flies different species? 5 primary forces affect genetic composition of populations and cause evolution • • • • • Natural selection Mutation Gene flow Genetic drift Mate choice Species • Species contain groups of interbreeding populations connected by gene flow. • The species is a natural and fundamental unit of evolution • Each species is the product of a unique and independent evolutionary pathway. How to define species • Biological concept – reproductively isolation • Phylogenetic concept – if a phylogenetic tree, each species uniquely occupies the tip of a tree branch • Morphological concept – consistent, distinguishing physical differences • Ecological concept How species form: Allopatric Sympatric Parapatric Founder Effect Three outcomes if hybridization With renewed or continued contact between two populations, there are three possible outcomes: 1. Individuals can hybridize readily. No speciation 2. Individuals do not hybridize at all. Full speciation 3. Individuals hybridize but offspring have reduced fitness. Speciation in progress. Selection for evolution of strong reproductive barriers. 46 How hybrids determine species • Are they reproductively isolated? • Can they interbreed back with parent groups? • Are they a separate but “weak” group that can interbreed? 1. Are they different species? 2. What species definition might apply here? 3. If they are (or are you thinking they might be), how did the speciation occur? Develop this idea. 4. Which evidences are the most important for your choice? Star those on your sheet. 5. What else do you need to know to validate your conclusion? Go online and research what you can find out. Think of a specific question you are trying to answer. (HW) Test Review • About Darwin • Evolution as defined by Darwin and concept today • Scientists/theories influenced development of the theory • Points of Darwin’s Theory • Evidences (4) for evolution • Homologous vs analogous structures • Vestigial structures • Microevolution vs macroevolution • Origins of genetic variation • Average heterozygosity • Hardy-Weinberg – how to do problems, conditions that maintain genetic equilibrium • How gene frequencies can be altered • Difference in types of genetic drift – bottleneck, founder effect • Relative fitness • Types of natural selection • Sexual selection – types • How to preserve genetic variation • 4 concepts of species • Types of reproductive isolation (prezygotic and postzygotic, types of each) • 3 main types of speciation, how differ • Autoploidy and allopolyploidy in sympatric speciation • Role of hybrids in determining speciation • Punctuated equilibrium vs gradual change in timing of evolution